Difference between revisions of "Use of Conifer Resin to Create a Hostile Environment for Pathogens in Wood Ant Colonies"

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==Introduction==
 
==Introduction==
Select a topic about genetics or evolution in a specific organism or ecosystem.<br>
 
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[[File:Wood ant (Formica lugubris) (33876593612).jpg |thumb|300px|right| Wood ants (Formica lugubris). Photograph taken by Gail Hampshire.]]
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[[File:Wood ant (Formica lugubris) (33876593612).jpg |thumb|300px|right| Fig. 1 Wood ants (<i> Formica Paralugubris </i>). Photograph taken by Gail Hampshire.]]
  
<br>At right is a sample image insertion. It works for any image uploaded anywhere to MicrobeWiki.  The insertion code consists of:
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As can be seen in Fig. 2, wood ants (<i> Formica paralugubris </i>) have been observed gathering pieces of solidified conifer resin and treating it with formic acid secretions for use as nest material <ref name = q>[https://doi.org/10.1111/j.1365-2311.2007.00983.x Castella G, Chapuisat M, Moret Y, Christe P. The presence of conifer resin decreases the use of the immune system in wood ants. Ecological Entomology. 2008 Jun;33(3):408-12.]</ref><ref name = c>[https://onlinelibrary.wiley.com/doi/abs/10.1002/ece3.2834 Brütsch T, Jaffuel G, Vallat A, Turlings TC, Chapuisat M. Wood ants produce a potent antimicrobial agent by applying formic acid on tree‐collected resin. Ecology and Evolution. 2017 Apr;7(7):2249-54.]</ref>
<br><b>Double brackets:</b> [[
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. This behavior can inhibit the growth of bacteria and fungi within the nest,  and can even protect wood ants against a variety of pathogens <ref name = q/>. Experiments testing the intentionality of this behavior have demonstrated that wood ants prefer to collect the resin over alternative building materials in the spring and summer months <ref name = q/>. This preference for resin seems to waver in autumn <ref name = q/>. No concrete answers have been given for this seasonal variation in behavior, but several hypotheses have been proposed to explain the phenomenon <ref name = q/>. One leading opinion is that the proportion of resin gathered drops as the ants focus on gathering twigs to prepare for the cold winter months <ref name = q/>. Another possibility is that because parasites can often be observed to time their reproduction with that of their host, the need for resin is much more significant in the spring and summer months, while the new brood of wood ant larvae is reared <ref name = q/>. Regardless of why this seasonal variation exists, the wood ants appear to recognize conifer resin through olfactory cues <ref name = q/>. The resin also appears to be used to prevent illness rather than treat it, as the proportion of resin gathered by infected ants is no higher than that gathered by uninfected ants <ref name = q/>. This behavior is a leading example of self medication behaviors in animals <ref name = q/>. Wood ants appear to gather the resin intentionally through olfactory cues, conifer resin is detrimental to a variety of wood ant pathogens, and this behavior does appear to have a positive effect on the fitness of wood ants in contact with pathogens <ref name = q/>.
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<br><b>Legend/credit:</b> Electron micrograph of the Ebola Zaire virus. This was the first photo ever taken of the virus, on 10/13/1976. By Dr. F.A. Murphy, now at U.C. Davis, then at the CDC.
 
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==Benefits of Conifer resin for wood ant health==
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[[File:Ant-e1492044072290.jpg |thumb|300px|left|Fig. 2 Wood ant carrying globule of conifer resin. Photograph taken by Arnaud Maeder]]
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The presence of conifer resin has a variety of demonstrable positive effects on the health, immune function, and survivability of wood ants. Immune function is an important  characteristic for protecting species from parasitic and pathogenic organisms <ref name = v> [https://core.ac.uk/download/pdf/18147903.pdf Castella G, Chapuisat M, Christe P. Prophylaxis with resin in wood ants. Animal Behaviour. 2008 Apr 1;75(4):1591-6.]</ref>. However, constant activation and maintenance of the immune system greatly reduces the fitness of most  organisms <ref name = v/>. This phenomenon has been directly observed in bumblebees, whose survivability under starvation conditions drops significantly with immune activation <ref name = v/>. Behavioral strategies for pathogen resistance, such as self-medication, are a useful biological strategy for mitigating these costs without compromising protection from pathogens <ref name = v/>. It is important to note that behavioral strategies are no more lacking in costly trade offs than traditional immune activities <ref name = v/>. These trade-offs can involve simple energetic costs, but may also come in the form of time lost for other beneficial tasks such as eating or reproducing <ref name = v/>. Still, behavioral strategies are often highly effective in increasing the overall fitness of an organism <ref name = v/>.
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<br><br>
  
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This principle can be demonstrated by the decrease in immune function that occurs when wood ants are in close proximity to conifer resin <ref name = v/>. Ants in resin-rich nests have been shown to have decreased antibacterial and lytic activities as compared to ants in resin-free environments <ref name = v/>. This decrease in immune function allows the ant metabolism to focus energy on beneficial metabolic pathways separate from immune function <ref name = v/>. This decrease also appears to have no detrimental effect on microbial resistance in the affected wood  ants, since the presence of conifer resin has also been shown to increase survival rates in colonies affected by a variety of pathogens <ref name = t> [https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2275180/ Chapuisat M, Oppliger A, Magliano P, Christe P. Wood ants use resin to protect themselves against pathogens. Proceedings of the Royal Society B: Biological Sciences. 2007 Aug 22;274(1621):2013-7.]</ref>. When infected with <i>Pseudomonas fluorescens</I> (soil bacteria which can have a highly detrimental effect on insects) both worker ants and larvae had significantly increased survival rates when nest material contained conifer resin <ref name = t/>. When infected with another pathogen, the parasitic fungus <i>Metarhizium anisopliae</i>, results were mixed <ref name = t/>. While conifer resin had little effect on the survival rate of adult worker ants exposed to <i>Metarhizium anisopliae</i>, it did appear to greatly increase the survival rate of ant larvae infected with the pathogen <ref name = t/>. It was also shown that the conifer resin had no positive or negative effects on ants in a pathogen free environment <ref name = t/>. This data is a testament to the strong beneficial effects incorporating conifer resin treated with formic acid into wood ant nesting materials has on wood ant pathogen resistance.
  
<br> <br>
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==Antimicrobial properties of Conifer resin==
 
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[[File:Conifer resin from Agathis lanceolata (New Caledonia, southwestern Pacific Basin) 5 (39352788962).jpg|thumb|300px|right|Fig. 3 Solidified conifer resin from New Caledonia. Photograph taken by James St. John]]
==Section 1 Benefits of Conifer resin for wood ant health==
 
Include some current research, with at least one image.<br><br>
 
[[File:Ant-e1492044072290.jpg |thumb|300px|left|Wood ant carrying globule of conifer resin. Photo taken by Arnaud Maeder]]
 
 
 
 
 
Sample citations: <ref>[http://www.plosbiology.org/article/fetchObject.action?uri=info%3Adoi%2F10.1371%2Fjournal.pbio.1000005&representation=PDF Hodgkin, J. and Partridge, F.A. "<i>Caenorhabditis elegans</i> meets microsporidia: the nematode killers from Paris." 2008. PLoS Biology 6:2634-2637.]</ref>
 
 
 
<ref>[http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3847443/ Bartlett et al.: Oncolytic viruses as therapeutic cancer vaccines. Molecular Cancer 2013 12:103.]</ref>
 
<br><br>A citation code consists of a hyperlinked reference within "ref" begin and end codes.
 
 
 
 
 
<ref>[https://doi.org/10.1111/j.1365-2311.2007.00983.x Castella G, Chapuisat M, Moret Y, Christe P. The presence of conifer resin decreases the use of the immune system in wood ants. Ecological Entomology. 2008 Jun;33(3):408-12.]</ref>
 
 
 
<ref>[https://doi.org/10.1111/j.1365-2311.2007.00983.x Castella G, Chapuisat M, Christe P. Prophylaxis with resin in wood ants. Animal Behaviour. 2008 Apr 1;75(4):1591-6.]</ref>
 
 
 
<ref>[https://doi.org/10.1111/j.1365-2311.2007.00983.x Chapuisat M, Oppliger A, Magliano P, Christe P. Wood ants use resin to protect themselves against pathogens. Proceedings of the Royal Society B: Biological Sciences. 2007 Aug 22;274(1621):2013-7.]</ref>
 
  
==Section 2 Antimicrobial properties of Conifer resin==
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Resin production and secretion is stimulated by mechanical wounding of a Conifer tree’s bark <ref name = f>[http://www.plantphysiol.org/content/129/3/1003 Martin D, Tholl D, Gershenzon J, Bohlmann J. Methyl jasmonate induces traumatic resin ducts, terpenoid resin biosynthesis, and terpenoid accumulation in developing xylem of Norway spruce stems. Plant physiology. 2002 Jul 1;129(3):1003-18.]</ref>. This system has evolved as a response to one of the greatest threats to conifer trees, bark beetle infestation <ref name = r>[https://pdf.sciencedirectassets.com/271203/1-s2.0-S1360138500X00349/1-s2.0-S1360138599014016/main.pdf?X-Amz-Security-Token=IQoJb3JpZ2luX2VjELP%2F%2F%2F%2F%2F%2F%2F%2F%2F%2FwEaCXVzLWVhc3QtMSJGMEQCIHMh%2B52o58eeq00xW%2FPuom2giQJJBk4GrWcAxOkJrB7CAiAspaS47dwE%2B%2B3M8i4OaKMivOT0sJT7Afpw4J4VDxtpoSq0AwhMEAMaDDA1OTAwMzU0Njg2NSIMcec5sEgE2oDCBa4rKpED%2F%2Bx6zg0HsOaZeDmB3wmxCdHRhrrYPKPiQ2XdR7xvX3reUxLAq1kpiFmANUPWviZsAzvgxLdik1ZMqPCl5%2BTDFkxUHh93FWuaIKmb3rDkElRl%2FQ2eYYcM9pFeQ3v5odDhZwblkz5wuRNB8%2BCPb%2BqeHQ6q%2FTyj6V%2FxtxxADtm2eFWmIltHRRGfMen3qcAdyNIb%2FklLThN2Qxw08KswtH1nkLA45X5t61oq7OI%2F7aRmmOkMUIsys9EB2ZY73qk7FGRJW%2B7zAQOOD4RFFx3fPLwVsbohBOifTOKcNni%2FyPNmgDm5CErdqSCzb1tTRy0upkECb9aic9ryHJt0yjlz83YvHGYRWTjYBOet28Hi7E%2FP%2FDTBibSg38UCYE1Btg1hTLN%2FQ9UbculxEHe02b%2BeSU487xWBOx2N9RkV6iAJxr7fqB5ZJ839g1wHoR7reZvR2teB9oicogDPRyFXAYVU18vM23k0QR2iUj3sYOdojaDICBs0DAUhQdJ0S%2BpeHWZ4c%2F%2FUCFND%2FudjL427thOfYzUZBJAwnY6%2F%2FgU67AFJSfLRBXkYs4UazHvwlB5RUmFhPXtBia4Hm%2FNxOIRnVcfHaNoIjPfOQDzly7Qs%2BpfHtxMJrAzW3UV2XNTfyyStoeqcH98ZyF59Kt%2BVG47e5w6yF%2FuB%2FFSLAi6L0iqJkAIy44U19JW%2BB7GNYTVCc6oCbmx%2BjGlemwIBs3gIiABjX8%2FeGRFCr3TYwE7TqCVlCzEcQTlfb6Qn9b7QSELaJsBOh5DogVr13JKkjelpX%2FLt%2F1mjAWhEMMhggU9t0a28fhc06JqxJsg2uvNvEgEW6axEODG%2F8j1l7P%2Fh54PWDNBulTXSAUxlVzFgLhZo%2FQ%3D%3D&X-Amz-Algorithm=AWS4-HMAC-SHA256&X-Amz-Date=20201208T194414Z&X-Amz-SignedHeaders=host&X-Amz-Expires=300&X-Amz-Credential=ASIAQ3PHCVTY3NNU2WBC%2F20201208%2Fus-east-1%2Fs3%2Faws4_request&X-Amz-Signature=ea2edbc0c03764e4772d3bd9f6fb2dad9179915e0090a13f312ae86df87fe922&hash=e17fce7b97a6829aed71c4a9b7d30e34887c6df5f939ee70b5715bfd8393e692&host=68042c943591013ac2b2430a89b270f6af2c76d8dfd086a07176afe7c76c2c61&pii=S1360138599014016&tid=spdf-e222bf30-c313-4e53-adf9-e6010a6032c5&sid=dcd514bf5c83d94b607a4809185b96d7670fgxrqa&type=client Phillips MA, Croteau RB. Resin-based defenses in conifers. Trends in plant science. 1999 May 1;4(5):184-90.]</ref>
Include some current research, with a second image.<br><br>
+
. The properties of resin, as displayed in Fig. 3, cause it to solidify when exposed to the open air <ref name = r/>. This trait allows resin to entrap the invading beetles, and is why the resin incorporated into ant nests is in the form of solid chunks <ref name = r/>. Beetles are not the only threat that comes with a bark beetle infestation however. The mechanical wounding of tree bark leaves conifers wide open to fungal infections <ref name = r/>. To deal with this problem, conifer resin contains a wide array of antimicrobial chemicals <ref name = f/>. These chemicals include limonene and 3-carene, as well as terpene based molecules including monoterpenes, sesquiterpenes and diterpenoid acid, all of which are highly toxic to bark beetles and their fungal co-conspirators <ref name = f/><ref name = r/>. These same terpene based molecules are used as olfactory cues by wood ants gathering conifer resin as a nest building material <ref name = q/>. These antimicrobial chemicals make conifer resin a highly effective tool for making a hostile environment for pathogens within wood ant nests <ref name = t/>. <I>Pseudomonas fluorescens</i> and <I>Metarhizium anisopliae</i>, two microbial agents mentioned above as wood ant pathogens, have their growth inhibited by the presence of conifer resin in vitro <ref name = t/>.This inhibition of microbial invaders is likely to extend to a greater number of wood ant pathogens in nature, especially with the addition of formic acid by ants. <ref name = c/><ref name = t/>.
[[File:Conifer resin from Agathis lanceolata (New Caledonia, southwestern Pacific Basin) 5 (39352788962).jpg|thumb|300px|right| Solidified conifer resin originating from New Caledonia. Photograph taken by James St. John]]
 
  
 
==Conclusion==
 
==Conclusion==
Overall text length should be at least 1,000 words (before counting references), with at least 2 images. Include at least 5 references under Reference section.<br><br>
+
The use of conifer resin to create a hostile environment for pathogens in wood ant colonies is a fascinating case of self-medication in non-human animals. This discovery has academic value on it’s own merits, but this research also sits on the cutting edge of inquiry into animal self-medication behaviors. While great strides have been made in the last few decades in understanding this seemingly strange wood ant behavior, there is still much to learn in this new and exciting field of study.
 
 
 
 
  
 
==References==
 
==References==

Latest revision as of 20:45, 9 December 2020

Introduction

Fig. 1 Wood ants ( Formica Paralugubris ). Photograph taken by Gail Hampshire.

As can be seen in Fig. 2, wood ants ( Formica paralugubris ) have been observed gathering pieces of solidified conifer resin and treating it with formic acid secretions for use as nest material [1][2] . This behavior can inhibit the growth of bacteria and fungi within the nest, and can even protect wood ants against a variety of pathogens [1]. Experiments testing the intentionality of this behavior have demonstrated that wood ants prefer to collect the resin over alternative building materials in the spring and summer months [1]. This preference for resin seems to waver in autumn [1]. No concrete answers have been given for this seasonal variation in behavior, but several hypotheses have been proposed to explain the phenomenon [1]. One leading opinion is that the proportion of resin gathered drops as the ants focus on gathering twigs to prepare for the cold winter months [1]. Another possibility is that because parasites can often be observed to time their reproduction with that of their host, the need for resin is much more significant in the spring and summer months, while the new brood of wood ant larvae is reared [1]. Regardless of why this seasonal variation exists, the wood ants appear to recognize conifer resin through olfactory cues [1]. The resin also appears to be used to prevent illness rather than treat it, as the proportion of resin gathered by infected ants is no higher than that gathered by uninfected ants [1]. This behavior is a leading example of self medication behaviors in animals [1]. Wood ants appear to gather the resin intentionally through olfactory cues, conifer resin is detrimental to a variety of wood ant pathogens, and this behavior does appear to have a positive effect on the fitness of wood ants in contact with pathogens [1].

Benefits of Conifer resin for wood ant health

Fig. 2 Wood ant carrying globule of conifer resin. Photograph taken by Arnaud Maeder

The presence of conifer resin has a variety of demonstrable positive effects on the health, immune function, and survivability of wood ants. Immune function is an important characteristic for protecting species from parasitic and pathogenic organisms [3]. However, constant activation and maintenance of the immune system greatly reduces the fitness of most organisms [3]. This phenomenon has been directly observed in bumblebees, whose survivability under starvation conditions drops significantly with immune activation [3]. Behavioral strategies for pathogen resistance, such as self-medication, are a useful biological strategy for mitigating these costs without compromising protection from pathogens [3]. It is important to note that behavioral strategies are no more lacking in costly trade offs than traditional immune activities [3]. These trade-offs can involve simple energetic costs, but may also come in the form of time lost for other beneficial tasks such as eating or reproducing [3]. Still, behavioral strategies are often highly effective in increasing the overall fitness of an organism [3].

This principle can be demonstrated by the decrease in immune function that occurs when wood ants are in close proximity to conifer resin [3]. Ants in resin-rich nests have been shown to have decreased antibacterial and lytic activities as compared to ants in resin-free environments [3]. This decrease in immune function allows the ant metabolism to focus energy on beneficial metabolic pathways separate from immune function [3]. This decrease also appears to have no detrimental effect on microbial resistance in the affected wood ants, since the presence of conifer resin has also been shown to increase survival rates in colonies affected by a variety of pathogens [4]. When infected with Pseudomonas fluorescens (soil bacteria which can have a highly detrimental effect on insects) both worker ants and larvae had significantly increased survival rates when nest material contained conifer resin [4]. When infected with another pathogen, the parasitic fungus Metarhizium anisopliae, results were mixed [4]. While conifer resin had little effect on the survival rate of adult worker ants exposed to Metarhizium anisopliae, it did appear to greatly increase the survival rate of ant larvae infected with the pathogen [4]. It was also shown that the conifer resin had no positive or negative effects on ants in a pathogen free environment [4]. This data is a testament to the strong beneficial effects incorporating conifer resin treated with formic acid into wood ant nesting materials has on wood ant pathogen resistance.

Antimicrobial properties of Conifer resin

Fig. 3 Solidified conifer resin from New Caledonia. Photograph taken by James St. John

Resin production and secretion is stimulated by mechanical wounding of a Conifer tree’s bark [5]. This system has evolved as a response to one of the greatest threats to conifer trees, bark beetle infestation [6] . The properties of resin, as displayed in Fig. 3, cause it to solidify when exposed to the open air [6]. This trait allows resin to entrap the invading beetles, and is why the resin incorporated into ant nests is in the form of solid chunks [6]. Beetles are not the only threat that comes with a bark beetle infestation however. The mechanical wounding of tree bark leaves conifers wide open to fungal infections [6]. To deal with this problem, conifer resin contains a wide array of antimicrobial chemicals [5]. These chemicals include limonene and 3-carene, as well as terpene based molecules including monoterpenes, sesquiterpenes and diterpenoid acid, all of which are highly toxic to bark beetles and their fungal co-conspirators [5][6]. These same terpene based molecules are used as olfactory cues by wood ants gathering conifer resin as a nest building material [1]. These antimicrobial chemicals make conifer resin a highly effective tool for making a hostile environment for pathogens within wood ant nests [4]. Pseudomonas fluorescens and Metarhizium anisopliae, two microbial agents mentioned above as wood ant pathogens, have their growth inhibited by the presence of conifer resin in vitro [4].This inhibition of microbial invaders is likely to extend to a greater number of wood ant pathogens in nature, especially with the addition of formic acid by ants. [2][4].

Conclusion

The use of conifer resin to create a hostile environment for pathogens in wood ant colonies is a fascinating case of self-medication in non-human animals. This discovery has academic value on it’s own merits, but this research also sits on the cutting edge of inquiry into animal self-medication behaviors. While great strides have been made in the last few decades in understanding this seemingly strange wood ant behavior, there is still much to learn in this new and exciting field of study.

References



Edited by Lucas Dunst, student of Joan Slonczewski for BIOL 116 Information in Living Systems, 2020, Kenyon College.